Exceptional crocodylomorph biodiversity of “La Cantalera” site ...

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Exceptional crocodylomorph biodiversity of “La Cantalera” site (lower Barremian; Lower Cretaceous) in Teruel, Spain

Eduardo Puértolas-Pascual, Raquel Rabal-Garcés, and José Ignacio Canudo

ABSTRACT

The palaeontological site of La Cantalera in Teruel, Spain (Blesa Formation,Lower Cretaceous) can be characterized as the site with the greatest biodiversity ofvertebrates (32 taxa) from the lower Barremian of the Iberian Peninsula. Remains ofamphibians, lizards, turtles, crocodiles, pterosaurs, dinosaurs and mammals havebeen recovered. The most common taxa are crocodylomorphs, and their most abun-dant remains are small isolated teeth. Six morphotypes have been identified, whichmay correspond to at least four different sympatric crocodylomorph taxa. This sitepresents a similar assemblage to other European Lower Cretaceous sites, with mor-photypes that may correspond to the families Goniopholididae, Bernissartiidae, Atopo-sauridae and multiple ziphodont crocodylomorphs. The crocodylomorph assemblageof La Cantalera is characterized by the absence of large individuals and presents awide variety of dental morphologies adapted to diets ranging from generalist to highlyspecialized. This rich biodiversity could be due to concentration of vertebrates intorestricted flooded areas during dry seasons within a marsh ecosystem. Furthermore,La Cantalera could also be close to a nesting area, which would explain the abundanceof small-sized individuals and the presence of fossil crocodiloid eggshells.

Eduardo Puértolas-Pascual. Grupo Aragosaurus-IUCA, Departamento de Ciencias de la Tierra, Facultad de Ciencias, Universidad de Zaragoza, Calle Pedro Cerbuna 12, 50009 Zaragoza (Spain). [email protected] Rabal-Garcés. Grupo Aragosaurus-IUCA, Departamento de Ciencias de la Tierra, Facultad de Ciencias, Universidad de Zaragoza, Calle Pedro Cerbuna 12, 50009 Zaragoza (Spain) [email protected]é Ignacio Canudo. Grupo Aragosaurus-IUCA, Departamento de Ciencias de la Tierra, Facultad de Ciencias, Universidad de Zaragoza, Calle Pedro Cerbuna 12, 50009 Zaragoza (Spain) [email protected]

Keywords: Cretaceous, Lower; Crocodylomorpha; teeth; palaeoecology; palaeobiodiversity; Iberian Range

PE Article Number: 18.2.28ACopyright: Society for Vertebrate Paleontology June 2015Submission: 2 October 2014. Acceptance: 22 May 2015

Puértolas-Pascual, Eduardo, Rabal-Garcés, Raquel, and Canudo, José Ignacio. 2015. Exceptional crocodylomorph biodiversity of “La Cantalera” site (lower Barremian; Lower Cretaceous) in Teruel, Spain. Palaeontologia Electronica 18.2.28A: 1-16palaeo-electronica.org/content/2015/1229-biodiversity-of-la-cantalera

PUÉRTOLAS-PASCUAL, RABAL-GARCÉS, & IGNACIO CANUDO: BIODIVERSITY OF “LA CANTALERA”

INTRODUCTION

There are several studies of palaeontologicalsites with vertebrate accumulations in the Barre-mian (Lower Cretaceous) of the Iberian Peninsula(Rauhut, 2002; Cuenca-Bescós and Canudo,2003; Ruiz-Omeñaca et al., 2004; Buscalioni et al.,2008; Canudo et al., 2002a, 2010). La Cantalera(Josa, Teruel) is of great interest because it is oneof the macro- and micro-vertebrate accumulationsites with the greatest palaeobiodiversity in theLower Cretaceous of Europe, with at least 32 dif-ferent taxa (Table 1) (Canudo et al., 2010). Thevertebrate assemblage is composed almost exclu-sively of terrestrial and amphibious organisms thatinhabited the vicinity of the site. La Cantalera hasbeen considered to provide a good temporal sam-ple of the faunal community that inhabited the area(Canudo et al., 2010; Gasca et al., 2014).

The most abundant skeletal remains found inLa Cantalera site are isolated teeth (Ruiz-Omeñaca et al., 1997; Canudo et al., 2004, 2010).The most common remains are crocodylomorphteeth. Together with disarticulated postcranialremains, these fossils testify to the high palaeobio-diversity of crocodylomorphs at this site. Teethbelonging to multituberculate mammals, periph-eral plates of turtles, cranial and postcranialremains of amphibians and lizards, isolated ptero-saur teeth and a single vertebra from an osteich-thyan fish have also been identified. The greatestbiodiversity at the site is associated with non-aviandinosaurs, with identified isolated teeth and post-cranial remains from: ornithopods, theropods, sau-ropods and thyreophorans (Ruíz-Omeñaca et al.,1997, 2001; Badiola et al., 2008; Canudo et al.,2010). Additional eggshell fossils have been recov-ered from the site, belonging to turtles, non-aviandinosaurs and crocodylomorphs (Moreno-Azanzaet al., 2009, 2014; Canudo et al., 2010). Further-more, coprolites with six different morphologieshave been identified (Ruíz-Omeñaca et al., 2001).

There have been relatively few studies ofcrocodylomorphs from the Lower Cretaceous ofthe Iberian Range (Buscalioni and Sanz, 1984,1987, 1990b; Buscalioni et al., 1984, 2008, 2013;Sanz et al., 1984; Brinkmann, 1992; Ortega et al.,1996; Ortega, 2004; Puértolas-Pascual et al.,2015). These studies have been undertaken overthe past three decades on the basis of isolatedremains and specimens represented by cranial andpostcranial material, primarily from the sites of Ter-uel (Josa, Galve, Ariño and Andorra) and Cuenca(Las Hoyas, Uña and Buenache de la Sierra).

Referring to Teruel, several studies of the Bar-remian have been carried out at palaeontologicalsites within the Galve syncline on the basis of teethand isolated postcranial and cranial fragments(Buscalioni and Sanz, 1984, 1987; Sanz et al.,1984; Ruíz-Omeñaca et al., 2004). In these papersthe crocodylomorph material was assigned toMesoeucrocodylia indet., Neosuchia indet., cf.Machimosaurus sp., Goniopholididae, Atoposauri-dae and Bernissartia. Furthermore, a completeskull attributed to Bernissartia fagesii Dollo, 1883was also described (Buscalioni et al., 1984; Bus-calioni and Sanz, 1990b).

In Josa, where the site of La Cantalera (lowerBarremian) is located, a preliminary study was car-ried out, focusing on a large sample of crocodylo-morph teeth. This allowed the teeth to be groupedinto different morphotypes, which were assigned tocf. Theriosuchus sp., Bernissartiidae indet., Gonio-pholididae indet. and cf. Lisboasaurus sp. (Canudoet al., 2010).

More recently, new studies have been carriedout in more modern sediments in the Escucha For-mation (lower Albian), close to the localities ofAndorra and Ariño. In these sites two new speciesof goniopholidids have been described (Alcalá etal., 2012; Puértolas-Pascual et al., 2012, 2015;Buscalioni et al., 2013; Puértolas-Pascual andCanudo, 2013).

There are also several other studies based onisolated teeth from the Lower Cretaceous ofEurope: the United Kingdom, France, Denmark,Sweden, Belgium and Portugal. Almost all of thesesites have a similar assemblage composed ofGoniopholididae, Bernissartiidae, Atoposauridaeand other less abundant forms (Buscalioni et al.,2008; Schwarz-Wings et al., 2009).

In this context, the aim of the present paper isto review and describe the crocodylomorph toothassemblage recovered from La Cantalera site inorder to determine the variability and biodiversity ofthe site and compare this with other European sitesof similar age.

GEOGRAPHICAL AND GEOLOGICAL SETTING

The palaeontological site of La Cantalera issituated in the northeast of the Iberian Peninsula,near the small locality of Josa in Teruel, Spain (Fig-ure 1.1).

Geologically, the site is located in the Meso-zoic Oliete Subbasin, which is one of the sevensubbasins belonging to the Late Cretaceous Mae-strazgo Basin (Soria de Miguel, 1997; Salas et al.,2001). This is located in the eastern part of the Ibe-

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PALAEO-ELECTRONICA.ORG

rian Range and borders the Tertiary basins of theEbro River in the north and the locality of Montal-bán in the south (Figure 1.1). During the Mesozoic,the Iberian Basin underwent a major extensionrelated to the opening of the Atlantic (Salas et al.,2001). The Oliete Subbasin was formed during the

second rifting stage that began in the Early Creta-ceous, when the carbonate platform of the centralsector of the Iberian Basin was fractured as aresult of the process of extension, creating severalsubbasins (Soria de Miguel, 1997; Salas et al.,2001).

TABLE 1. Faunal assemblage of La Cantalera. Colors show the ecological affinity of each taxon, aquatic in blue,amphibious in green and terrestrial in brown. Modified from Canudo et al., 2010 and Ruiz-Omeñaca et al., 1997.

Class Order/Superorder Sub/Infraorder Taxa

Charophyceae Charales Quinquespiralia Atopochara trivolvis triquetra Grambast, 1968

Pseudoglobator sp.

Gastropoda Caenogastropoda Viviparidae indet.

Basomatophora Basomatophora indet.

Ostracoda Podocopida Podocopina Cypridea (Ulwellia) soriana Kneuper-Haack, 1966

Osteichtyes Teleostei indet.

Lissamphibia Albanerpetonidae indet.

Anura Discoglossidae indet.

“Reptilia” Chelonia Pleurosternidae? indet.

Crocodylomorpha Goniopholididae?

Bernissartiidae?

Theriosuchus sp.

Mesoeucrocodylia indet.

Squamata Lacertilia Paramacellodidae? indet.

Pterosauria Istiodactylidae? indet.

Ornithocheiridae? indet.

Ornithischia Ornithopoda Iguanodontoidea indet.

Hadrosauroidea? indet.

“Hypsilophodontidae” indet.

Rhabdodontidae? indet.

Thyreophora Polacanthidae indet.

Saurischia Sauropoda Euhelopodidae indet

Theropoda Baryonychinae indet.

Carcharodontosauridae? indet.

aff. Paronychodon sp.

Maniraptora indet. 1



Dromaeosaurinae indet. 1



Velociraptorinae indet.

Avialae? indet.

Mammalia Multituberculata Cantalera abadi Badiola, Canudo and Cuenca-Bescós, 2008

Eobaatar sp.

“Plagiaulacida” indet.

Plagiaulacidae/Eobaataridae indet.

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The Lower Cretaceous of the Oliete Subbasinis characterized by four megasequences: the Mar-gas y Calizas de Blesa Formation; the Calizas deAlacón Formation; the Margas de Forcall Forma-tion; and the Calizas de Oliete Formation (Soria deMiguel, 1997) (Figure 1.2-4). The site of La Can-talera forms part of the typical “Wealden facies” ofthe Lower Cretaceous. More specifically, it islocated within the gray clays of the lower part of theBlesa Formation (Figure 2), which has traditionallybeen considered upper Hauterivian / lower Barre-mian in age (Soria et al., 1995; Canudo et al.,2002b; Aurell et al., 2004) due to the presence ofthe charophyte Atopochara trivolvis triquetraGrambast, 1968. However, the presence of thisspecies at the base of the Blesa Formation indi-cates a lower Barremian age, as in the upper partof the Castellar Formation at Galve (Canudo et al.,2012).

In the area around Josa, the Blesa Formationdisplays a complex sedimentary evolution. Conti-nental sedimentation predominates at the bottom,with alluvial, lacustrine and palustrine deposits(Figure 2), while the top shows facies of marshesand coastal lagoons. The site of La Cantalera hasbeen interpreted as mudflat deposits associatedwith swampy areas with periodic droughts in anarea of marshy vegetation. The significant concen-tration of vertebrate remains at the site of La Can-talera can perhaps be explained by the small sizeand isolated nature of the swampy domains, whichmight have attracted animals for feeding (Ruíz-Omeñaca et al., 1997; Aurell et al., 2004).

MATERIALS AND METHODS

The materials included in this study wererecovered by washing and sieving (sieves of 2, 1and 0.5 mm mesh) over three tons of sediment

FIGURE 1. Geographical and geological setting of La Cantalera site (early Barremian, Teruel, Spain). 1, Geologicalmap of the Iberian Peninsula with detailed location of the site of La Cantalera and its position within the MaestrazgoBasin (Ol: Oliete, Pa: Las Parras, Ga: Galve, Mo: Morella, Pe: Perelló, Sa: Salzedella, Pg: Peñagolosa). 2, Geologicalmap of the area of La Cantalera. 3, Local geological units. 4, Stratigraphic column and position of La Cantalera. Mod-ified from Canudo et al., 2010.

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recovered during several field campaigns from1997-2000. The study includes a total of 365 croc-odylomorph teeth, usually with a size between 1and 2 mm in height, although some specimenssmaller than 0.5 mm or around 1 cm in height werealso found. Broken or very worn teeth have notbeen taken into account in the measurements table(see Appendix).The figured and measured speci-mens have been designated with the abbreviationsMPZ (Museum of Natural Sciences of the Univer-sity of Zaragoza). All teeth were examined underan Olympus SZ40 binocular microscope with anLC20 digital camera incorporated and a scanningelectron microscope SEM JEOL JSM 6360 (20kV,Working Distance = 15mm) at the University ofZaragoza. The picture scaling and measurementswere performed with the software Olympus LCmi-cro (developed by Olympus Soft Imaging Solu-tions) and ImageJ (rsb.info.nih.gov/).

The fossil vertebrate assemblage from the LaCantalera site described in this paper is housed inthe Museo de Ciencias Naturales de la Universi-dad de Zaragoza (MPZ).

SYSTEMATIC PALAEONTOLOGY

Because of the great intraspecific variation intooth morphology and a shortage of phylogenetic

information, it is difficult to assign a taxonomic sta-tus at generic and even at family level to isolatedteeth belonging to Crocodylomorpha (Prasad andBroin, 2002; Buscalioni et al., 2008). We have thusdecided to be conservative with the taxonomicassignment of each morphotype.

Superorder CROCODYLOMORPHA Walker, 1970 (sensu Clark, 1986)

Clade MESOEUCROCODYLIA Whetstone and Whybrow, 1983 (sensu Benton and Clark, 1988)

Clade NEOSUCHIA? Benton and Clark, 1988Family GONIOPHOLIDIDAE? Cope, 1875

Genus and species undeterminedFigure 3

Referred material. 169 isolated teeth (46.30% ofthe sample = 365 teeth; MPZ 2014/468, 469, 470,471a, 471b and 719 through 820).Description. The referred teeth are representedby two different sizes, with large teeth rangingbetween 5 and 10 mm (Figure 3.1–2), and verysmall sizes ranging between 1 and 2 mm (Figure3.3–4). They are conical in shape, with generalistforms having a circular cross-section, with mediumto high crowns and a very acute apex that isslightly lingually curved. The lingual and labial sur-faces are separated by smooth carinae, the labialsurface being slightly more convex than the lingual

FIGURE 2. General view of La Cantalera site, showing the gray clays of the lower part of the Blesa Formation.

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face. The carinae show different degrees of devel-opment, being more marked in the smaller teeth(Figure 3.4). The transition between the tooth androot is almost unconstricted. In radicular view, thepulp cavity is short and shows subcircular morphol-ogy. Enamel ornamentation is formed by betweenfive and 10 longitudinal and almost parallel ridgesgoing straight from the base and convergingslightly toward the apex. In some teeth, the longitu-dinal ridges next to the carinae of the lingual faceseem to diverge.Remarks. The dental morphologies describedabove are often attributed to Goniopholididae. Nev-ertheless, very similar teeth are also present inmost neosuchians with generalist habits and diets,and they are also common in the anterior dentalregion of several crocodylomorphs with heterodontdentition, such as Bernissartia fagesii (Buffetautand Ford, 1979) or Theriosuchus (Schwarz andSalisbury, 2005; Lauprasert et al., 2011; Martin et

al., 2014). However, the high percentage of thisform of tooth (46.30% of the sample) is unlikely tohave come from only teeth from the anterior posi-tions of heterodont crocodylomorphs, so there islikely at least one taxon present at La Cantalerawith a generalist dentition, such as Goniopholidi-dae.

Family BERNISSARTIIDAE? Dollo, 1883Genus and species undetermined

Morphotype AFigure 4.1-3

Referred material. 104 isolated teeth (28.49% ofthe sample = 365 teeth; MPZ 2014/472, 473, 474a,474b and 843 through 901).Description. Molariform or tribodont teeth, bulky,bulbous, rounded, with low crowns and a bluntapex. The teeth have circular, elliptical (Figure 4.2),kidney-shaped (Figure 4.3) or figure-eight-shapedcross-sections at their base (Figure 4.1). Certainspecimens display stronger mediolateral compres-

FIGURE 3. Crocodylomorph teeth attributed to Goniopholididae? from La Cantalera site. 1–3, MPZ 2014/468, 469and 470; photographs taken with a binocular microscope in mesial/distal, labial, lingual, apical (above) and radicular(below) views. 4, photographs taken with a scanning electron microscope in labial (above, MPZ 2014/471a) andmesial/distal (below, MPZ 2014/471b) views. Scale bars equal 1 mm.

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sion than others (Figure 4.1), but they all show arounded profile in lingual/labial view. The base ofthe crowns has a strong constriction. The pulp cav-ity, which can only be seen in a very few speci-mens because the root is not usually preserved,presents sub-ellipsoidal morphology, with the majoraxis running anteroposteriorly. Enamel ornamenta-tion is formed by thin parallel ridges convergingtoward the apex. In most teeth, the presence ofmesiodistal carinae is unlikely. However, in tallerand less worn teeth a basiapical keel that is moremarked towards the apex can be observed.

Within this morphotype there is some degreeof variability according to the heterodont dentitionobserved in complete skulls and jaws of Bernissar-tia (Buffetaut and Ford, 1979; Buscalioni and Sanz,1984, 1990b). The teeth from the posterior regionof the tooth row have lower crowns, kidney-shapedor figure-eight-shaped sections and greater lateralcompression (Figure 4.1, 3). By contrast, the ante-riorly placed teeth have morphologies with tallercrowns and sub-ellipsoidal to circular sections,without figure-eight-shaped sections (Figure 4.1).Almost all teeth have a worn occlusal surface.

FIGURE 4. 1–3, MPZ 2014/472, 473, 474a and 474b; crocodylomorph teeth from La Cantalera site attributed to Ber-nissartiidae?; photographs taken with a binocular microscope (1,2) in mesial/distal, labial, lingual, apical (above) andradicular (below) views; photographs taken with a scanning electron microscope (3) in lingual (MPZ 2014/474a,above) and apical (MPZ 2014/474b, below) views. 4–6, MPZ 2014/475, 476, 477a, 477b and 477c; crocodylomorphteeth from La Cantalera site attributed to Bernissartiidae?, which may correspond to teeth from the anterior or midregion of the tooth row; photographs taken with a binocular microscope (4,5) in mesial/distal, labial, lingual, apical(above) and radicular (below) views; photographs taken with a scanning electron microscope (6) in labial (MPZ 2014/477a, left), mesial/distal (MPZ 2014/477b, right) and apical (MPZ 2014/477c, below) views. Scale bars equal 1 mm.

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Remarks. This morphotype has been assigned toBernissartiidae? since this type of morphology iscommon in taxa of the same age, such as Bernis-sartia or Koumpiodontosuchus (Buffetaut and Ford,1979; Sweetman et al., 2014). However, care mustbe taken with such specific assignments. This typeof tooth, associated with a durophagous diet, isalso present in other taxa not closely related withBernissartiidae, such as Unasuchus, Acynodon,Allognathosuchus, Brachychampsa, Alberto-champsa and the extant Osteolaemus (Buffetautand Ford, 1979; Brinkmann, 1992; Buscalioni etal., 1997, 1999; Martin, 2007; Delfino et al., 2008).Therefore, the presence of this type of morphologymay be more closely linked to dietary function thanto phylogenetic relatedness (Buffetaut and Ford,1979).

Morphotype BFigure 4.4–6

Referred material. 62 isolated teeth (16.98% ofthe sample = 365 teeth; MPZ 2014/475, 476, 477a,477b, 477c and 902 through 962).Description. These teeth are short and conicalwith a triangular outline in lingual/labial view andstrong mediolateral compression. The crowns arerelatively low and small in size (1–2 mm). Thelabial surface is convex, and the lingual face isslightly flattened. The apex is slightly linguallycurved, and there are mesial and distal carinae.The root contour is somewhat constricted, and thepulp cavity has subcircular to ellipsoidal morphol-ogy that is elongate in an anteroposterior direction.On the lingual surface, the base of the tooth has aslight concavity, giving it a somewhat kidney-shaped cross-section at its base. The ornamenta-tion consists of thin parallel longitudinal ridges thatconverge towards the apex on both sides.Remarks. In previous papers, such teeth havebeen attributed to forms close to Lisboasaurus andthe LH-Crocodyliform (crocodyliform from LasHoyas, Cuenca, Spain) (Canudo et al., 2010)which are two mesoeucrocodylians from the Kim-meridgian / Barremian of Spain and Portugal (Bus-calioni et al., 1996; Ortega, 2004; Schwarz andFechner, 2004, 2008). The triangular profile withlaterally compressed crowns and the presence of alingual groove at the base of the crown are indeedreminiscent of teeth present in Lisboasaurus andthe LH-Crocodyliform. However, the teeth from Lis-boasaurus and the LH-Crocodyliform have com-pletely smooth enamel, lacking carinae andornamentation (Schwarz and Fechner, 2008). Theornamentation of this dental morphotype from LaCantalera and its constriction at the base of the

crown are also consistent with teeth attributed toBernissartiidae. The greater crown height and themore conical shape could correspond to teeth fromanterior positions in the jaw or to unworn teeth fromthe mid region in the dental series of Bernissartii-dae (Buffetaut and Ford, 1979; Buscalioni andSanz, 1990b; Sweetman et al., 2014).

Family ATOPOSAURIDAE Gervais, 1871Genus THERIOSUCHUS Owen, 1879

Species undeterminedFigure 5

Referred material. 26 isolated teeth (7.12% of thesample = 365 teeth; MPZ 2014/478, 479, 480, 481,482a, 482b, 482c and 821 through 841).Description. The teeth show the typical autapo-morphic characters observed in the genus Therio-

suchus, with dentition that comprises labio-lingually compressed low-crowned, pseudocanini-form and lanceolate-shaped teeth, with mesial anddistal carinae. The teeth are laterally compressedwith differences in size. The labial surface is con-vex and flatter than the lingual face, with a slightcurvature towards the lingual area near the apex.This morphotype shows morphological variabilitythat can be related to different dental positionsobserved in skulls and jaws with preserved teeth ofTheriosuchus (Owen, 1879; Brinkmann, 1992;Schwarz and Salisbury, 2005; Karl et al., 2006;Lauprasert et al., 2011; Martin et al., 2014). Theteeth located in posterior positions of the tooth row(Figure 5.1–2, 5) are wider mesiodistally, highly lat-erally compressed, leaf-shaped and with a lowercrown. The labial surfaces of these teeth are con-vex, and the lingual surfaces are flatter in the baseand somewhat concave towards the apex. In addi-tion, the radicular contour is slightly constricted,and the pulp cavity is completely flattened andelongated. By contrast, the teeth from anteriorpositions (Figure 5.3–4) are less laterally com-pressed, are lanceolate and have a considerablyhigher crown. They have mesial and distal carinae,but their torsion towards the lingual area is lesspronounced. The pulp cavity is not as elongated orflattened, presenting an ellipsoidal morphology.Remarks. Within this morphotype there existsgreat morphological variability associated with theenamel ornamentation and the presence orabsence of true or false denticles on the carinae.Most teeth present longitudinal well-marked ridges,which are convergent in the central area of thetooth and diverge towards the carinae (Figure 5.1,3, 5); others have fine anastomosed striae (Figure5.2), and yet others have almost smooth enamel

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(Figure 5.4). This kind of tooth has been associ-ated with insectivorous diets (Buscalioni and Sanz,1988). The presence, absence and size of the den-ticles on the carinae seem to be related with theconvergence of enamel ridges or grooves towardsthe carinae (“false ziphodont” according to Prasadand Broin, 2002). The size of the denticles are pro-portional to the thickness of the ridges. However,teeth with nearly smooth enamel may also havedenticles (Figure 5.4). The presence of leaf-shapedor lanceolate teeth is probably related to the toothposition in the dental series, but the variation in theenamel ornamentation may be due to the presenceof several taxa or great intraspecific variability. Thiskind of dental morphology is typical of atoposauridssuch as Theriosuchus.


Family, Genus and species undeterminedFigure 6.1

Referred material. 3 isolated teeth (0.82% of thesample = 365 teeth; MPZ 2014/483 and 842).Description. Conical teeth with high crowns, a cir-cular cross-section at the base and a sharp apexcurved lingually and distally. Their relatively largesize varies between 5 and 10 mm. The enamelornamentation is somewhat variable and may besmooth or with fine longitudinal ridges. The lingualand labial surfaces are separated by carinae withtrue denticles (ziphodont), with the labial surfaceslightly more convex than the lingual face. A toothis ziphodont when the denticles are not the resultof the prolongation of the enamel ridges (Prasadand Broin, 2002).Remarks. The term “ziphodont” has long beenapplied to Mesoeucrocodylia, including several

FIGURE 5. Crocodylomorph teeth attributed to Theriosuchus sp. from La Cantalera site. 1–4, MPZ 2014/478, 479,480 and 481; photographs taken with a binocular microscope in mesial/distal, labial, lingual, apical (above) andradicular (below) views. 5, MPZ 2014/482a, 2014/482b and 2014/482c; photographs taken with a scanning electronmicroscope in mesial/distal (2014/482a, above), lingual (2014/482a, middle) and apical (2014/482a, below) views.Scale bars equal 1 mm.

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genera from such a broad range of families thatthey may well not be monophyletic. Typical taxawith true ziphodont dentition are notosuchians, pla-nocraniids and some peirosaurids. The use ofziphodont dentition as a criterion for the taxonomicclassification of isolated teeth must thus be takenwith caution since it is considered to be of limitedvalue as phylogenetic information (Turner, 2006;Andrade and Bertini, 2008). This type of dentitionwas present in highly predatory terrestrial crocodyl-omorphs, thus having ecological rather than phylo-genetic implications (Andrade and Bertini, 2008).For this reason, we have chosen to assign thismorphotype to Mesoeucrocodylia indet.


Family, Genus and species undeterminedFigure 6.2

Material. 1 isolated tooth (0.27% of the sample =365 teeth; MPZ 2014/484).Description. Relatively large tooth (about 1 cm)with spatulate morphology, mediolaterally com-pressed and with a low crown. The lingual andlabial faces have two longitudinal grooves dividingthe enamel into three bulbous surfaces that aremore pronounced towards the base of the tooth.The enamel is smooth and has mesial and distalcarinae with true denticles of variable size. One ofthe carinae forks into two at the base of the tooth.Remarks. The overall tooth morphology resemblesthat of the spatulate teeth of Atoposauridae. How-ever, we do not know other crocodylomorphs withthis kind of enamel ornamentation and the pres-ence of bifurcated carinae, so it has been assignedto Mesoeucrocodylia indet. Given that only onetooth with this strange morphology has been found,

FIGURE 6. 1, MPZ 2014/483; ziphodont crocodylomorph tooth attributed to Mesoeucrocodylia indet. in mesial/distal,labial, lingual (with detail of the denticles), apical (above) and radicular (below) views. 2, MPZ 2014/484; bizarre croc-odylomorph tooth attributed to Mesoeucrocodylia indet. in mesial/distal (arrows show the bifurcated keel), labial, lin-gual, apical (above) and radicular (below) views. All photographs taken with a binocular microscope. Scale bars equal1 mm.

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the carinae could be the result of some form ofpathology.

PALAEOECOLOGY

The palaeontological site of La Cantalera wasformed in a small basin, with no fluvial influence orallochthonous accumulation of sediments, andwhere most of the recovered fossils show no evi-dence of transport. La Cantalera has been inter-preted as comprising floodplain deposits withephemeral lacustrine episodes in an area ofmarshy vegetation (Aurell et al., 2004). The tapho-nomic, stratigraphic and sedimentological charac-teristics of La Cantalera indicate that the highconcentration and biodiversity of vertebrates at thesite is the result of sediment accumulation in a len-tic ecosystem and not an accumulation caused bytransport. It is therefore a site with a very concretespatio-temporal distribution that shows the faunathat inhabited the area for a relatively short timeinterval (Canudo et al., 2010), and thus it is reason-able to assume sympatry of the species.

Given the faunal assemblage of a palaeonto-logical site, it is possible to make predictions andinterpretations about the type of ecosystem thatoccupied the area at the time (Buscalioni et al.,2008). The taxa of La Cantalera were divided intodifferent groups (aquatic, amphibious and terres-trial) (Table 1) according to their dependence onwater for extant wetlands (Valk, 2006; Buscalioni etal., 2008; Gasca et al., 2012). Gasca et al. (2012)proposed a simplified model based on Valk (2006),considering as aquatic taxa to organisms that can-not survive without the presence of water, such asfishes or aquatic algae. Amphibious taxa would bethe species that spend part of their life cycle inwater and part on terrestrial environments but theycannot survive if the wetland is totally drained, agood example would be the lissamphibians orsome turtles and crocodiles. Terrestrial species areorganisms that can be found both in wetlands andterrestrial environments and they can survive out-side the wetland, such as lacertids, pterosaurs,dinosaurs and mammals. The results show that16.21% of the organisms were aquatic, with twospecies of charophytes, two of gastropods, one ofostracod and just one osteichthyan fish. Amphibi-ous organisms show a similar percentage at18.91%; these are composed of at least four croco-dylomorphs, two lissamphibians and one turtle(very scarce). Terrestrial organisms show the high-est percentage at 64.86%, and these are made upof one lizard, two pterosaurs, 17 dinosaurs andfour mammals.

The large number of terrestrial organisms inrelation to the aquatic and amphibious fauna con-trasts with the association expected in a wetlandecosystem, where aquatic and amphibious speciesare often the most abundant (Valk, 2006), as atother sites of similar age such as Buenache or LasHoyas (Cuenca, Spain) (Buscalioni et al., 2008).The site of La Cantalera shows a faunal assem-blage more similar to the site of Uña (Cuenca).However, the large number of terrestrial organismsin Uña is attributed to taphonomic and sedimento-logical factors, since the abundance of theseorganisms can be explained by transport andallochthonous sedimentation (Buscalioni et al.,2008). The scarce transport observed in La Can-talera, together with other factors such as theabundance of terrestrial organisms and the highnumber of replacement teeth of herbivorous dino-saurs, indicates that La Cantalera could have beenan occasional feeding area for herbivorous dino-saurs, especially during wet periods (Ruíz-Omeñaca et al., 1997).

Another remarkable thing in La Cantalera isthe small size of the recovered remains. Except forsome large bones belonging to dinosaurs, most ofthe remains are small teeth. This also applies tothe crocodylomorphs, with tooth heights ofbetween 1 and 2 mm on average, and only a fewspecimens are less than 0.5 mm or around 1 cm inheight. 52.60 % of the crocodylomorph teeth corre-spond to Bernissartiidae? and Atoposauridae.These taxa are characterized by their small bodysize, with a total length of around 50 cm in the adultstage and never exceeds 1 m in length (Buscalioniand Sanz, 1990a, 1990b, 1990c; Schwarz-Wingset al., 2009). Despite their small body size, theteeth of adult atoposaurids and bernissartiids werebigger than those from La Cantalera (average sizeof 1.79 mm for Theriosuchus sp. and 1.31 mm forBernissartiidae?, see Appendix) and thus most ofthe teeth from La Cantalera likely belong to juve-niles. For example, the teeth from adult specimensof Theriosuchus pusillus Owen, 1879 or Theriosu-chus sympiestodon Martin, Rabi, Csiki-Sava andVasile, 2014 range between 2 mm (posterior maxil-lary teeth) to 10 mm (enlarged fourth maxillarytooth) in height, while young individuals rangebetween 1 mm (posterior maxillary teeth) to 3.5mm (enlarged fourth maxillary tooth) in height; andthe teeth from adult specimens of bernissartiidssuch as Koumpiodontosuchus or the holotype ofBernissartia from Belgium range between 3 mm to6 mm in height (measurements taken from the fig-ures of Owen (1879), Clark (1986) and Martin et al.

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(2014) for Theriosuchus; from Sweetman et al.(2014) for Koumpiodontosuchus; and firsthandmeasurements taken by us for Bernissartia).Therefore, the teeth from adult specimens of Theri-osuchus or Bernissatiidae are above the 2 mm inheight, while in juvenile specimens are usually lessthan 2 mm in height, with the exception of the 4thenlarged maxillary tooth which is usually greaterthan 2 mm in height. Given the small size of mostof the teeth from La Cantalera attributed to Goniop-holididae? (average size = 2.28 mm, see Appen-dix), the presence of adult goniopholidids is ruledout, since this is a group that can reach up to 4 mof body length with teeth with an average size of 13mm in the adult stage (Buscalioni et al., 2008). Thisargument would be strengthened because imma-ture specimens of dwarf goniopholidids such asNannosuchus gracilidens Owen, 1879 have teeththat vary between 2.5 to 7 mm in height (measure-ments taken from the plate 44 of Owen (1879)),which is the approximate size range observed in LaCantalera. The high concentration of probablejuvenile crocodylomorphs and the abundant croco-dylomorph eggshell fragments (Canudo et al.,2010; Moreno-Azanza et al., 2013) at the site of LaCantalera could be explained by the presence of anearby nesting area. Extant crocodylomorphs, thehatchlings and juveniles usually remain around thenesting area and nearby areas with the mother forweeks or even until two seasons before dispersing;freshwater wetland areas/grasslands are the mostcommon places for nesting (Brazaities and Wata-nabe, 2011).

CONCLUSIONS

La Cantalera has at least six morphotypesthat could correspond to at least four different croc-odylomorph taxa (Goniopholididae?, Bernissartii-dae?, Theriosuchus and an undeterminedziphodont). Due to the high variability within eachmorphotype, especially those assigned to Therio-suchus, the presence of more than four differentcrocodylomorph species seems the most likelyhypothesis.

The site has a similar crocodylomorph assem-blage to other European Lower Cretaceous sitessuch as Bornholm (Denmark) and Buenache de laSierra (Cuenca), where there is also an associationof Goniopholididae, Bernissartiidae, Atoposauridaeand an undetermined ziphodont (Buscalioni et al.,2008; Schwarz-Wings et al., 2009). We prefer notto assign the bizarre tooth with bifurcated carinaeto any particular taxon, because there is only one

isolated specimen with this morphology and itcould be the result of pathology.

This site is also characterized by the absenceof large individuals and by the presence of teethwith a wide variety of morphologies. Such toothdisparity has been explained in terms of a trophicdiversity ranging from generalist to insectivorousand durophagous habits (Buscalioni and Sanz,1988). Furthermore, this rich biodiversity could bedue to the fact that in marsh ecosystems, espe-cially during dry periods, vertebrates tend tooccupy and move to areas that remain flooded,where water and food are more abundant. Anotherhypothesis that could explain the abundance ofsmall teeth and the presence of crocodylomorpheggshell fragments is that La Cantalera was occa-sionally used as a nesting area

ACKNOWLEDGEMENTS

This work forms part of the EuropeanRegional Development Fund, the Government ofAragón ("Grupos Consolidados" and "DirecciónGeneral de Patrimonio Cultural") and the programFormación del Profesorado Universitario (FPU)subsidized by the Spanish Ministry of Education,Culture and Sports. The comments of AlexanderHastings have improved the original manuscript.Rupert Glasgow edited the text in English.

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APPENDIX - TEETH MEASUREMENTS

Appendix material is available online at palaeo-electronica.org/content/2015/1229-biodiversity-of-la-cantalera

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Exceptional crocodylomorph biodiversity of “La Cantalera” site ...

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